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Marine Biology 90, 379-394 (1986

)

Marine

:===BiOlOgy
© Springer-Verlag 1986

Synchronous spawnings of 105 scleractinian coral species on the Great Barrier Reef
R. C. Babcock, G. D. Bull, P. L. Harrison, A. J. Heyward, J. K. Oliver, C. C. Wallace and B. L. Willis Department of Marine Biology, James Cook University; Townsville, Queensland 4811, Australia

Abstract

Following observations of mass spawning of hermatypic corals on the Great Barrier Reef in 1981 and 1982, spawning dates were successfully predicted and documented at five reefs on the Central and Northern Great Barrier Reef in 1983. During the predicted times, 105 species from 36 genera and 11 families were observed to spawn. Of these, 15 species were shown to have an annual gametogenic cycle. All but two of the species observed during mass spawnings shed gametes which underwent external fertilization and development. Synchronous spawning was observed both within and between the five reefs studied, which were separated by as much as 5 ° of latitude (500 kin) or almost a quarter of the length of the Great Barrier Reef. The mass spawning of corals took place on only a few nights of the year, between the full and lastquarter moon in late spring. Maturation of gametes coincided with rapidly rising spring sea temperatures. Lunar and diel cycles may provide cues for the synchronization of gamete release in these species. The hour and night on which the greatest number of species and individuals spawned coincided with low-amplitude tides. Multispecific synchronous spawning, or "mass spawning", of scleractinian and some alcyonacean corals represents a phenomenon which is, so far, unique in both marine and terrestrial communities.
Introduction

Kojis and Quinn (1982b) suggested that the broadcasting of gametes for external fertilization during a brief annual spawning was probably the most common mode of reproduction among scleractinians. This was confirmed by Harrison et al. (1984), who showed that more species of coral are known to broadcast gametes than to brood their offspring. Harrison etal. (1984) documented spawning in corals on reefs in the central Great Barrier Reef region, and showed that the majority of these broadcasting species spawned together on the same nights. These spawning

events were synchronous in that individual colonies and species spawned during only a few hours over several consecutive days every year. Other studies of scleractinian reproduction had pointed to a distinct seasonality of spawning on the Great Barrier Reef(GBR) (Bothwell, 1982; Harriott, 1983); nevertheless, the fact that populations of many widespread and abundant corals spawned together on only the same few lunar nights every year had not previously been documented. There are a number of well known examples of highly predictable, highly synchronized annual reproductive cycles in marine organisms. Spawning may occur on only one or two nights a year (e.g. in Eunice viridis, Korringa, 1957; Comanthusjaponica, Kubota, 1981) or it may occur with a definite periodicity within a more extended breeding season [Centrechinus (Diaderna) setosa, Fox, 1924; Leuresthes tenuis, Korringa, 1957; Neofibularia nolitangere, Reiswig, 1970; Clunio rnarinus, Neumann, 1981]. These instances all represent spawning events restricted to a single species at a particular time and place. Although a number of species of hydromedusae have been reported to spawn together at Friday Harbour, USA (Miller, 1979), the spawning occurs over an extended period. The mass spawning of corals differs from all the above phenomena in the synchronous spawning of many species in a very brief period of time. This paper presents findings from an ongoing program to document the mode and timing of coral spawning, and to more fully document the numbers of species involved in the mass spawning of corals on the GBR. Other objectives were to determine whether the mass spawning occurred at reefs in the northern as well as the central region, and to establish the relative timing of spawning in the two regions.
Materials and methods

Study sites Coral spawning was studied at four sites in the central GBR: Big Broadhurst Reef, Bowden Reef, Magnetic Island

Goniastrea aspera. A.e. A. BBR: Big Broadhurst Reef. Observations of coral spawning were also made at Lizard Island in the northern region of the GBR (Fig. A. At night. LI: Lizard Island. G. in order to determine whether part of the population delayed spawning until a later period.380 PAPU A R. A. 25 of which spawned within one day of each other (Table 8). Babcock et al. hyacinthus (3). A. 1984). showing study areas. G. millepora and Galaxea fascicularis were taken every month between July and November 1983 at Orpheus Island. A. Fifty species were recorded spawning at more than one reef (Table 1). Spawning activity was also inferred by several indirect means. and corals in aquaria. Montipora digita and Platygyra sinensis were sampled more intensively during the spawning period (i. Magnetic Island is situated in the shallow (< 10 m) inshore waters of Cleveland Bay.. A. BR: Bowden Reef. flulchra (2). Scleractinians participating in the spawning event included 36 genera in 11 families.: Synchronous spawnings of scleractinian corals and the presence of testes (Kojis and Quinn. 20 species contained mature gonads when sampled during the first few days of the spawning period (Table 2) but for logistic reasons could not be sampled further. These were the presence of pigmentation in the eggs A total of 105 species of scleractinian corals and 7 species of alcyonacean corals. mieroflhthalma (1). while all the other sites are mid-shelf reefs or islands in deeper ( > 20 m) offshore waters. Babcock. colonies were broken open and examined in the field or brought back to the laboratory and checked under a dissecting microscope. A. the presence of eggs or sperm in aquaria.. latistella (2).. in order to monitor gametogenic cycles. Fig. five randomly selected colonies of Goniastrea aspera. These species were chosen since they were already the subjects of separate detailed reproductive studies. and what proportion of the population spawned on each day of the spawning period.C. A. nobilis (2). To determine whether gonads were in a ripe condition. This included direct observations of spawning in 33 species. Results Taxonomic and geographic extent of mass spawning Gametogenic cycles Fifteen species of coral were sampled throughout the year at Magnetic Island and Orpheus Island. 1981. or the presence of eggs in plankton mesh bags placed over colonies in the field. Mass spawning therefore involved whole populations of many species of coral on widely separated reefs. 15 or more colonies of each. In addition to the 105 species which were observed to spawn. valida (2). ~' '. Two criteria were used to determine whether gonads were ripe. Motile sperm with condensed heads indicated that spawning would take place within the week (Harrison et al. Map of the Queensland coast. A. three to five random samples of A. Acropora formosa. Frequent brief observations were made using flashlights. In addition. OI Orpheus Island and Orpheus Island. . corals in aquaria were shielded as much as possible from artificial illumination. favulus. from five geographically distinct reefs. Microscopic examination of smears of squashed fresh testes was used as a final measure of gonad maturity. formosa (7). These species were Acropora elseyi (1 colony). MI: Magnetic Island. 1984. millepora (4). Gamete release was observed directly in 80 of the 107 scleractinian species. 1. Coral spawning was observed on the same nights at reefs in the central and northern Great Barrier Reef region separated by up to 500 kin. At Magnetic Island. In the remaining species gamete shedding was inferred from the abrupt disappearance of gametes between sampling periods and the absence of brooded embryos or planulae. corals moved to underwater observation posts.' r' i: Spawning records Direct observations of spawning were made at night on undisturbed corals in situ. including the disappearance of gonads between sequential samples from tagged colonies in the field or in aquaria. Two planulating species of coral were also recorded. °. tenuis (1). humilis (2). released gametes during a seven-day spawning period (Table 1). 1984). At both sites. 1). favulus and Platygyra sinensis were sampled at monthly intervals between March and November 1983. tagged colonies of 11 species of Acropora were sampled at approximately monthly intervals between January and October 1983.. Harrison et al.. Tables 3 and 4).

6 (BR) 6 2:10 2:05. lamellosa Sperm in aquaria Planulae in coelenteron Planulae released yeHow MI MI D D 14 4-7 LI. BBR MI. BBR MI. 5 (OI). white white pink pink pink. red red pink red cream. grandis A. tenuis A. horrida E. BBR) 3:10 (LI). 3-?-7 (Oi) 2 1:25 4. BR MI. LI MI. BBR: Big Broadhurst Reef. spumosa M. 6 (BR) 6 0:10 (MI). 6. gemmifera A. except in cases where indirect spawning records are the only ones available. Tubastrea faulkneri Faviidae A ustralogyra zelli Barabattoia amicorum Caulastrea furcata Cyphastrea chalcidicum C. 3:50 (BR)s. LI. valida A. A. formosa A. 6 (LI). 5. florida A. secale A. 6. 5 (LI). 3:50 (BR)s - I I I I I H H H 3 (MI). 5 (LI). BR BBR MI. 3:10 (BR). 4:05 (BBR) 1-?-7 (MI). hispida M. BR: Bowden Reef. 2:40 (BBR) 1:50 (MI). MI: Magnetic Island.R. 2:05 (LI) 3:50 (BR)s - I I I I I I I I I pink pink pink. vigorous ejection o f gametes. pir~k orange. h:min) Type of spawning behaviour Egg colour Scleractinia Acroporidae A cropora aculeus A. 2 (OI) 2 4 1. 2:10. cytherea A. 7 (LI) 3 (MI). BR. 6(BBR) 6 3 (MI). release not observed. 6 (BBR) 1. 5 (OI) 5. latistella lutkeni Iongicyathus loripes microphthalma millepora BR BBR OI. OI OI MI. 3:45 (LI) 3:40 (LI) 3:40 (BR) I I I I I I I I H H H 7 5. Babcock et al. OI H H 5 6 2:00 - I I H H 4 (OI). Type III. s: setting o f egg-sperm bundles only. turgescens Agaricidae Pachyseris rugosa P. 2 2 H H D D D D 5 6 4. D: dioecious. 16 6 2:30 (MI) 3:00 2:45 (OI). LI. humilis A. 6 (BR) 3 6 2:20s 3:00 2:35 (MI). LI.: not recorded Species Site of observations Sexual character Day of spawning (days after full moon) Hour of spawning (hours after sunset. LI: Lizard Island. monasteriata M. nobilis A. 5. yongei Astreopora myriophthalma Montipora digitata M. 5 ( B R ) l:55 (MI). 1. pink cream pink red white. 5 (MI). pulchra A. 1:15. 5. 2:25 (BR) - H H H H H H 4 (LI). purple tan (zooxanthellate) tan (zooxanthellate) tan tan (zooxanthellate) tan (zooxanthellate) tan pink yellow yellow orange H H H H H H H H A. LI. C. passive release o f gametes. austera A. foliosa M. . 7 (LI) 3 (MI). 2 (MI). 2:10 (OI) 2:25 2:40 2:25 2:15 2:50 2:25 I:10 (OI) 0:10 (MI). 5. BR MI. BR BBR OI. OI LI. 3:40 (OI). 4. 2:45 (MI). pink cream. OI: O r p h e u s Island. 0:40 (OI) 3:30 1:25 I I I I I I I I II II Caryophyllidae Euphyllia divisa OI Physogyra lichtensteini OI Dendrophyllidae Dendrophyllia sp. selago A. 3:15 (LI) 2:30 (MI). 6 (BR) 6 (BR) H H H H H 5 (BR) 3 (MI). 7 (LI). tuberculosa M. 1:40 (LI) 3:50 (BR)s 2:50 (BR) 1:55 (MI). 2:25 (MI). 3:40 (BR). BR BR. 4 (OI). OI MI OI MI MI OI BR OI. 1:00 (OI). 6 (O1) 5 4 2:05 (LI). BR MI BBR LI BR. 3:15. LI. Data for day and hour o f spawning are from direct observation only. OI. 6. 3:10. LI. 7 (MI). A. microphthalma Echinopora gemmacea E. Type II. red red pink. cerealis A. 1:45 ( B R ) pink green pink pink grey-brown pink pink (continued overleaf) H H H H 5 4. 5. nasuta A. hyacinthus A. LI. A. 7 (LI) 4 (OI). 4 (OI). 6 (BR) 15. LI. BBR LI MI. speciosa 3 6 3 3 (MI). robusta A. 2:05 - 1:05 (OI). 1:05 (LI) 2:25 (MI). 6 (LI) 3(MI). 1:25. H: hermaphroditic. 3:40 (LI). List o f corals recorded spawning in 1983. OI MI. BR MI BBR MI LI MI MI. A. elseyi A. OI MI. sarmentosa A. Spawning behaviour: Type I. 3:50 (BR)s - I I m H H H H 15(MI). 6 (BR) 6 3:40 (OI). LI BR MI.: Synchronous spawnings o f scleractinian corals 381 Table 1. BR LI. 6 (BR). A. 6 (BR). 6(BR). BR MI OI LI MI H U H H 5 (LI. slow extrusion o f gametes. 5 6 6 2:35 1:40 3:50 s 1:40 .

LI) 6 6 (MI). OI. 3:40 (OI) 2:00 <2:10 0:40 0:40 (OI). hemprichii Symphyllia radians S. brown aqua. palauensis G. 2:55 (OI.. BBR) 4. 2:10 (BBR) 4:20 (LI)s 1:50 (MI). OI) 1-?-6 2-?-8 5 6 5 4. 5 (OI) 1:10. 5 (LI).. OI 1982) H H H H H H H H D D D 6 5 (OI. LI H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H 5 (LI) 6 (BBR) 2-?-6 4 (OI). BBR MI.382 Table I (continued) Species S i t e of observations R. h:min) 1:30 (LI) 1:50 (OI). 5 (LI) 3-? 6 2-?-8 2:05 3:25 (BBR). green pink pink pink pink. BBR. pectinata G. 3:25 (MI). 5 (LI. red red cream. BR. LI MI. 5 (OI). OI MI. 5 (MI) 6 3. complanata F. 1:10. BBR MI. OI LI. 4 (OI). 5 (OI) 5 (BBR) 5-?-6 2-?-6 4 (MI). 7 (BBR) 4 (OI) 5. BBR LI MI. BR Mussidae Acanthastrea eehinata Lobophyllia eorymbosa L. 7. 5 (OI). LI LI. 5 (LI) 4 (MI). 6 (BR). pink pink yellow pink. LI MI. lamellina P. 1:55 (OI). 1:20 (LI). yellow I I pink pink pink brown brown brown II . blue lavender cream. 3:10. 4:30 (LI) 1:20 (MI) 1:35 3:25 (MI). OI MI MI. faseieularis MI. LI. 2:25 (BR). 4 (MI). BBR Oxypora glabra LI O.5 4. 5. 4. sinensis Fungiidae Fungia fungites Sandalolitha robusta OI. 4 (LI). stelligera F. LI. 6 (OI) 5 (BBR) 3. pentagona E russelli Goniastrea aspera G. lactuca BBR P. 1: 15 (MI). 6 (0I). BBR LI. C. LI G. 4. 6 (OI). bennettae F. 2:55 (LI). OI.5 5 (LI) 6 (MI). rigida Leptoriaphrygia Montastrea curta M. LI) 2:40 3:10 (OI). 3:10 (oI) MI BBR D D H H H H 5 5 6 5 (LI). 1:40 (OI) 3:45. 3:10 (OI) 3:00-3:50 (BBR) l: 10 (MI). LI MI. lizardensis F. 4. 8 (MI) 3-?-6 5 (OI. 6. OI LI. BBR MI. orpheensis OI. BBR) 6 (MI). veroni Favites abdita F. brown red grey pink. OI 1:30 (LI). valenciennesi Moseleya latistellata OulophflIlia crispa Platygyra daedalea P. OI OI OI. 1:15 (LI) yellow yellow pink. 4 (MI). 4. LI. Babcock et al. lacera OI Pectinia alcicornis OI P. 4. BBR BBR MI. 5:00 (LI) 3:35 2:20 (OI). 1:35 (LI) 1:25 (MI). LI BBR. 2:50 1:55. OI. 3:40 (BR) 2:40 (OI) 1 : 15. red pink pink pink 0:55. LI MI. 1:10 (OI). 6 (BBR). pink _Pectiniidae Echinophyllia aspera MI E. BR. 2:10 (LI) 1:15 (MI). pallida F. 5 (OI) 5 (MI) 5 (BR) 4. 0:40 (OI) 1:40 (BBR) I II H H H H H 2:20 (MI). 6 (LI) 3. 2:40 (BR) I 2-?-8 (oi) 4. 5. 1:25 0:55. red pink yellow red pink. LI LI. OI OI. 4:10 (LI) - Type of spawning behaviour II II II I I I I I II I II I I II lII I I I II II II (LI). 5. matthai F. pini P. 6 (BR) . BBR BBR MI MI. 3:20 (OI). minor MI Goniopora sp. red aqua. 5. 5 (LI) aqua. LI. OI OI. 5 (LI. BBR OI OI OI.4 6 4 (OI). magnistellata M. retiformis Hydnophora exesa H. some sperm jets separate II l 1 pink white. 5 (OI) 2-?-8 4. OI BR. 3:25 (LI). red. LI LI OI. 6 (LI) 2-?-8 6 (MI). 5 (OI. (OI) 3. -0:05 (OI) I-II. 5. BBR OI MI. 1 (Veron and Pichon. recta Oculinidae Galaxea astreata G. OI.favulus G. OI MI MI LI. 3:40 (OI) 3:25 (BBR). BBR Mycedium elephantotus MI. 4 (MI). OI. LI) 6. 6. edwardsi G. LI. 5 (BR).: Synchronous spawnings of scleractinian corals Sexual character Day of spawning (days after full moon) Hour of spawning (hours after sunset. 5. I (O1) I I I I Egg colour Faviafavus F. LI) 1:55 (MI) 3:40 (OI. 6 (BBR) 5 (BBR).0:05 3:30 Ejected sperm every 4 min I creamy-yellow creamy-yellow pink apricot pink tan orange Merulinidae Clavarina triangularis OI MeruRna ampliata LI. LI. paeonia OI Poritidae Goniopora lobata OI. halicora F. flexuosa F. 0:25 (OI) 2:15 (MI) 2:55 (BR) 2:20. red pink green. 4. 5 (LI). 4. red tan tan.

hirsutum Sarcophyton n. -: not recorded Species Site of observation No. BBR: Big Broadhurst Reef. LI: Lizard Island. E g g c o l o u r w a s v a r i a b l e . of colonies Sexual character Last day sampled Egg colour Acroporidae A cropora anthoceris A. I m m a t u r e o o c y t e s w e r e first v i s i b l e i n d i s s e c t e d s a m p l e s i n late s u m m e r ( J a n u a r y . sp. solida Alcyonacea Alcyoniidae BR OI OI MI MI D D D D D tan A lcyonium aspiculatum Lobophytum compactum L. OI: Orpheus Island. lutea P. Galaxea fas- cicularis. P i n k or r e d was . BBR OI LI OI BBR LI LI LI 1 5 4 5 4 1 3 1 1 1 1 1 2 1 1 1 1 1 1 2 H tt H H H H H H H H H D D D D D H D D D 6 6 6 6 6 6 6 1 3 6 6 (OI) 6 5 3 2 6 purplebrown pink pink white orange tan brown - Favites chinensis Montastrea annuligera Fungiidae Heliofungia aetiniformis Herpetoglossa simplex Herpolitha limax Podabaeia crustaeea Polyphyllia talpina Mussidae Lobophyllia paehysepta Siderastreidae Coseinarea eolumna Thamnasteriidae Psammoeora eontigua P.R. T h e eggs o f t h e f a v i i d species b e c a m e pigm e n t e d f o u r to six w e e k s p r i o r to t h e s p r i n g s p a w n i n g . Babcock et al.f.J u l y ) i n o t h e r species. valeneiennesi Astreopora sp.. favulus. Porites cylindrica P.5 4 - 4 4 4 4. G o n a d m a t u r a t i o n c o n t i n u e d u n t i l eggs w e r e s h e d i n s p r i n g ( O c t o b e r o r N o v e m b e r ) . lobata P. polydactyla OI OI OI OI OI Ol OI 4 -- -- 4. a n d t h o s e o f Acropora species a n d Galaxeafascicularis a p p r o x i m a t e l y t h r e e w e e k s b e f o r e t h e i r release. G. clathrata A.5 0:40 - I - red Table 2. deformis S. but for which no subsequent observations or samples could be made. H: hermaphroditic. 1 (Veron) Montipora turtlensis Faviidae BBR BBR BBR BBR BBR BBR BBR LI BBR LI BBR LI OI. divaricata A. 1 Sinularia c. "Epidemic spawning" Diffuse sperm clouds - Egg colour Goniopora sp. Coral species with mature gonads immediately prior to major spawning period. a n d in m i d .w i n t e r ( J u n e . Goniastrea aspera. digitata G a m e t o g e n i c cycles A n n u a l g a m e t o g e n i c cycles w e r e o b s e r v e d in all species s a m p l e d t h r o u g h o u t t h e y e a r (Aeropora spp. Platygyra sinensis). samoensis A. h:min) 1:40 2:35 sperm. 2:45 eggs 2:40 ( 383 Type of spawning behaviour II II. monticulosa A. D: dioecious. e v e n w i t h i n species ( T a b l e 1). "Last day sampled" refers to n u m b e r of days after full moon.: Synchronous spawnings of scleractinian corals Table 1 (continued) Species Site of observations Sexual character Day of spawning (days after full moon) 5 4 5 3-?-7 3-?-7 Hour of spawning (hours after sunset. subulata A. microlobulatum L.F e b r u a r y ) i n A cropora species. C.

~ ~Z o~ F © II I I I I II II o m © o~ m I II I I I I II II . 0 ~Z I.: Synchronous spawnings ofscleractinian corals ..-~ 0 ~ .~ ~ 0 6 [ " o © oq 0"~ ~ .. Babcock et al.384 R.=.~ 0 ~ 0.a # .. o ~ "0 .~'~ ~ o ~ ~ o Z "0 ~ 's.9 ~ & / . "0 r _L • ~ ~= .'o~ o © I ]- ~ ~ .~ ~ ~ ~'.. C..~ .o dm O ~o~ ..~ O I II I I I I II II © ~d " ~0 © r~ .o ~ ~' ~d I l r I I I I I I I I .. = =~= .~0 "q .

~ lb ld I ld I ld . 2d lb 2a . la 2b 3b 2b 3b -. 5d (1) r 2f. 2 d I 2d p a r t i a l . 2b. full m o o n w a s 20 N o v e m b e r . - - If 3b 2a la lb la 4a - - o. 2d la. 4b.la. 4c 24Nov.~. formosa A. m 43c - la 2a. M o n t i p o r a digitata Caulastrea f u t r a t a A cropora cerealis A. T o t a l n u m b e r o f species o b s e r v e d = 4 8 . r e t i f o r m i s P l a t y g y r a lamellina Lobophyllia hemprichii Galaxea fascicularis F a v i t e s halicora Goniastrea aspera P l a t y g y r a sinensis M o n t i p o r a tuberculosa Leptoria phrygia Mycedium elephantotus P e c t i n i a alcicornis P. _ 26Nov.. ~ N ~ ~ ~_~ ~ .I ~ eq ~ t'N 2a 3b 3a la 2b la 2b . _ 25Nov. 50c 4a 23Nov... B a b c o c k et aL: S y n c h r o n o u s s p a w n i n g s o f s c l e r a c t i n i a n corals 385 Table 4. 2b . ~ ~ ~. 1 0 d . p e c t i n a t a A canthastrea e c h i n a t a O x y p o r a laeera Goniopora sp. 2d . O r p h e u s I s l a n d coral s p a w n i n g r e c o r d s . 2a. C. o f c o l o n i e s o b s e r v e d on: 21.2a-~ la.~ ~ ~ ~ b~ . 2b - 3b 5a 2b 4b la 2b. b e n n e t t a e Favites flexuosa Goniastrea p a l a u e n s i s G. o f species for w h i c h direct s p a w n i n g o b s e r v a t i o n s w e r e m a d e = 3 8 . 1 ± 50 4 1 3 3 2 1 1 1 4 3 5 5 3 4 1 4 11 9 15 7 3 3 11 6 16 18 1 1 1 6 4 1 5 2 1 4 3 2 1 1 1 1 1 2 1 2 22 N o v . no. millepora A.. l d .~. l d lb.R.2b 4a.Nov. nasuta M o n t i p o r a hispida P h y s o g y r a lichtensteini O u l o p h y l l i a crispa Porites cylindrica A eropora tenuis P a c h y s e r i s rugosa Montastrea magnistellata Lobophyllia corymbosa G o n i o p o r a lobata Favites abdita Echinopora gemmacea Favia pallida Favites complanata Goniastrea f a v u l u s G. 4b. 2d .. 4d 4a. 1983.5a.~.I ld 1 2dl ld -I 2d -I ld t "" . d [ 3a.~ I t I I la t 3a._S ~. - 1 - la la lb. F o r f u r t h e r details see l e g e n d to T a b l e 3 Species n N o . 2a. 2b 2a..I o. 3d 10d I 2a. p e o n i a Porites lobata P a e h y s e r i s speciosa P l a t y g y r a daedalea E u p h y l l i a divisa Platygvra pini E e h i n o p h y l 6 a orpheensis Astreopora myriophthalma Clavarina triangularis F a v i a veroni F. 2b.~ 4b 10d I 4b lb la lb 3a.lb-~ 2a.

Table 8). A.. The time of release was also consistent from year to year. Although populations of most coral species at each site spawned during only one spawning period. yet at the end of the spawning period the colony still contained significant numbers of gametes...... and 3 h 25 min after sunset in 1983 (Table 8). testes containing spermatids were observed only in the four to six weeks prior to the October or November spawning. Some species showed a range of spawning times within a population (e. but on the fifth and sixth nights after the full moon at the three other reefs (Tables 3-7). Goniastrea aspera. Acropora latistella spawned just after new moon at Magnetic Island (Table 3). nasuta.. ~6 ' . .. . Q1415 AFTER FULL i6 MOON . In a further 22 species... Galaxea fascicularis. ..g. 1984).. .. and Platygyra sinensis).. For example. at least five colonies were recorded spawning during one to three consecutive nights on one reef (Tables 3-7).. Montastrea valenciennesi.. both in terms of the number of species and the number of colonies observed to spawn. Bottom abscissa gives date of spawning. Hour of spawning Release of gametes in the synchronous multispecific spawning event began around sunset and continued for approximately four hours (Table 1). Favites pentagona. Number of coral species observed spawning on each day is given. all colonies sampled from a single reef were found to release gametes over one to three consecutive nights (Tables 3 and 4). Tables 3-7). At least 87 of the 105 species recorded spawned on these nights (Table 1. at Magnetic Island Acropora tenuis spawned 20 min after sunset in 1982 (Harrison etal.386 most common.. In 17 of the 33 species observed at more than one reef. an early inshore spawning in October at Magnetic Island. Only a few species were recorded to spawn on the first and second nights.. In dissections of fresh specimens collected throughout the year. intermittent abscissa. humilis. OCT MONTH NOV Fig.. Babcock et al.. 2. filled circles: new moon . Acropora formosa and A. and in some cases pigment was absent and eggs appeared white. Another type of split spawning period was apparent within one colony of Galaxea fascicularis (Table 4) which was observed spawning in the field. R. Fig. astreata... Motile sperm were not detected in freshly prepared squashes until approximately one week prior to spawning. one species was observed to have a sprit spawning period which took place in two consecutive months (A cropora latistella)..C. A.. Montipora digitata.. G. Astreopora myriophthalma. these were largely benthic and immobile. A similarly divided spawning period was inferred in other species from the absence of gametes in part of the population prior to the observed spawning at the site. Barabattoia amieorum. Only those species for which the exact day of spawning could be determined have been included. 45 40 35 30 25 20 15 10 5 O1~4ss78 DAYS . and 10rain after sunset in 1983 (Table 8). ~6 . Mobile planulae were found in the coelenteron of Dendrophyllia sp.g. showing days after full moon. Favia pallida... G. In contrast.. Platygyra sinensis spawned 3 h 40 rain after sunset in 1982 (Babcock. or seventh and eighth nights after full moon. Colonies of Tubastrea faulkneri released large (2 mm 10ng) bright orange planulae in aquaria over a few days. Sequential samples of the same two colonies over a period of 22 d SO. Table 8) and between populations (e.. and a later offshore spawning exactly one lunar month afterwards in November at the other four reefs. favulus. but spawned after the full moon at Big Broadhurst Reef (Table 6).. elseyi also spawned after the new moon at Magnetic Island (Table 3)... Peak spawning occurred on the third to the fifth nights after the full moon at Magnetic and Orpheus Islands. A greatly extended spawning period was observed in Hydnophora exesa which released eggs singly in aquaria over several nights at Magnetic Island (Table 3). populations spawned within an hour of each other on the same lunar day (Table 8). . The time of spawning (hours after sunset) was generally consistent within each population and between populations at different sites. Galaxea fascicularis. Of the corals studied only two species were observed to release or contain planulae. although colours ranged from brown to aqua. tenuis.. . A.. Lunar day of spawning Two discrete multispecific spawning periods were recorded in the spring of 1983. 36 . .. 2o . 2)." Synchronous spawnings ofscleractinian corals Spawning synchrony within populations In the five detailed population studies (involving Acropora formosa. or from the presence of maturing gametes in a few colonies some days after the rest of the population had spawned (Acropora millepora.. open circles: full moon. the third to tile sixth nights after the full moon were the major nights of spawning activity. At all five reefs. unpublished data). has been drawn only for those days on which sampling or observations were carfled out (see Tables 3-7). Frequencies of coral spawnings in 1983.

Echinophyllia aspera) or in a r e g u l a r cyclic fashion (Fungia fungites). 1983. Histological section (5/zm) of freshly spawned egg-sperm bundle. 3. nasuta A. The p a c k a g i n g process took 20 to 60 rain to be completed. o: oocytes. 1 Merulina ampliata Acropora nobilis A. full moon was 20 November. Type I spawning behavior. e: egg-sperm bundle. humilis A. We have t e r m e d this stage of the process the "setting" o f the egg-sperm b u n d l e . G a m e t e s were ejected in a variety of forms: as egg-sperm b u n d l e s (G. Goniastrea palauensis. A n i n t e r m e d i a t e s p a w n i n g b e h a v i o u r (Types I . Fig. formosa A. of species for which direct spawning observations were made = 19. immediately following the release of egg-sperm bundle. 387 Acropora aculeus A.).R. 25 Nov. Table 1. millepora A. of colonies observed on: 22-24 Nov.. .I I ) was observed i n Galaxeafascicularis. (Aquarium photo) showed a progressive decrease in the n u m b e r s of gametes at each sampling. I n h e r m a p h r o ditic colonies the eggs a n d s p e r m w i t h i n each polyp were p a c k a g e d into o n e or m o r e c o m p a c t egg-sperm b u n d l e s (Type I. aspera) or separate- 0-5 mm Fig. Bowden Reef coral spawning records. Videotape recordings o f a n A cropora elseyi c o l o n y s p a w n i n g showed the egg-sperm b u n d l e b e i n g rotated u n d e r the d i s t e n d e d oral disc o f each polyp. Lobophyllia hemprichii). Total no. For further details see legend to Table 3 Species n No. 4. s: sperm ly as eggs a n d s p e r m clouds (G. gemmifera A. cerealis A. Spawning behaviour Three types o f s p a w n i n g b e h a v i o u r were observed in corals participating in mass s p a w n i n g (Table 1). R o t a t i o n o f the b u n d l e c o n t i n u e d d u r i n g the final release phase as the polyp m o u t h o p e n e d a n d the b u n d l e was slowly released from the polyp. d u r i n g which time the b u n d l e was visible in the p h a r y n x or u n d e r the d i s t e n d e d oral disc of each polyp. The most comm o n type o f s p a w n i n g b e h a v i o u r (Type I. Table 1) was the slow release o f g a m e t e b u n d l e s t h r o u g h the p o l y p m o u t h which took place t h r o u g h o u t the c o l o n y over a p e r i o d o f 5 to 50 m i n (Acropora spp. hyacinthus A.: Synchronous spawnings of scleractinian corals Table 5. Babcock et al. favulus. This process was repeated once or several times i n quick succession (Goniastrea aspera. Platygyra spp. which slowly released egg-sperm b u n d l e s together with separate jets of sperm. secale A. grandis Cyl~hastrea chalcidicum Goniastrea pectinata PIatygyra daedalea Goniopora sp. I n Type II s p a w n i n g b e h a v i o u r . Platygyra sinensis. of species observed=19. 26 Nov. Note that none of the oocytes have been fertilized. no. selago A. 3). valida Montipora turgescens 2 1 1 2 1 1 8 8 5 5 2 1 1 2 2 1 3 1 1 - la la 1a 2a Ia la 3a 3a - 5a 5a 5a 5a 2a la la 2a 2a la 3a la la Fig. C. the gametes were ejected from the polyp via r a p i d contractions over all or part o f the colony after a short p e r i o d o f setting.

of colonies observed on: 23Nov. lb )b la I 1 lb lb if lb lb. austera A. no. tenuis A. No sign of fertilization was observed prior to the fragmentation of the egg-sperm bundles (Fig. but appeared to drift passively out from the gaping mouth of each polyp. These spawning observations encompass about one-third of the 340 .5 h after spawning. The spawning of Goniastrea f a v u l u s was unique among the corals observed in that the eggs were contained in a sticky mucous envelope and were negatively buoyant. hyacinthus A. Total no. The eggs and egg-sperm bundles of most gametespawning corals species. valida 1 1 1 1 3 1 2 3 2 2 2 3 1 1 1 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 24Nov. Sandalolitha robusta A ustralogyra zelli Favia stelligera Favites halicora Goniastrea edwardsi Echinophyllia orpheensis A streopora myriophthalma Platygyra daedalea Mycedium elephantotus Symphyllia radians Favia lizardensis Platygyra pini Favia pallida Montastrea curta Pectinia lactuca Favites abdita Goniastrea ?ectinata Lep toria phrygia Symphyllia recta A cropora secale A. ld ~ld -2d - 26Nov. Discussion Our results. After the eggsperm bundles reached the surface they broke apart. of species observed=37. and the larvae did not become strongly mobile until approximately 36 h old.: Synchronous spawnings of scleractinian corals 6. robusta A. of species for which direct spawning observations were made=34. 25Nov.388 Table R. The first signs of fertilization were not observed until approximately 2. lb lb la lb 3a lb lb t la la. whether hermaphroditic or dioecious. florida A. cytherea A. gametes were neither squeezed out or forcibly ejected from the polyp. ld-< lb---q lb lb I I I t--ld I ] - 2d 2d lb lb lb lb lb lb - - lb lb lb lb lb lb lb lb lb lb lb lb A third type of spawning beliaviour (Type III. nasuta A. lutkeni A. full moon was 20 November. elseyi A.C. We have now documented the mode and timing of reproduction for species belonging to 11 of the 15 families ofhermatypic scleractinians on the GBR. loripes A. including those of Harrison et al. 1983. The buoyant nature of the reproductive products of the majority of species probably increased the rate of fertilization by concentrating gametes at the surface. The eggs of Astreopora myriophthalma also appeared to sink after separating from buoyant egg-sperm bundles. humilis A. were buoyant and floated to the surface layers of the sea. gemmifera A. Big Broadhurst Reef coral spawning records. yongei A. millepora A. (1984) show that at least 109 species of scleractinians and 7 species of soft corals broadcast gametes in multispecific spawning episodes. Table 1) was apparent in Hydnophora exesa. 27Nov. In this species. longicyathus A. 4). For further details see legend to Table 3 Species n No. cerealis A. re- leasing the eggs and sperm. Babcock et al.

ld 3d ld-3d 2d lb. gemmifera A. Babcock et al.lb~ 2b lb. ld. d.. R a n d o m s a m p l e s t a k e n at o t h e r times o f the y e a r f r o m a wide range o f coral species failed to detect r i p e n i n g o f gonads on this massive scale in these or o t h e r species not r e c o r d e d in this paper. 1981..ld.t lb. 1933. d.- F I I I I . Wallace. b. l d .t 3b. 2 d . 1983.. 26 Nov. predictability a n d brevity o f the mass coral s p a w n i n g a p p e a r to be l i n k e d to successive e n v i r o n m e n t a l . 3d lb. C. H e y w a r d a n d Collins. (partial) -------4 lb. of colonies observed on: 22 Nov. Total no. loripes E c h i n o p o r a horrida P a c h y s e r i s rugosa Hydnophora exesa H. p i n i P.lb-I lb. - 24 Nov. 1"--t 2b lb. 6* lb-t 2 d . 3d.lb--t l d . p a l l i d a Favites flexuosa Goniastrea e d w a r d s i G. lb lb lb --lb I 25 Nov. Kojis a n d Q u i n n . Robertson. Lizard Island coral spawning records. 1985. secale 1 5 1 4 1 2 1 1 1 1 1 1 2 2 9 1 2 2 3 1 3 2 3 2 2 1 1 3 9 2 1 4 2 5 5 1 1 1 1 4 2 3 t I J I I - 389 23 Nov. sinensis Merulina ampliata Loboph/llia hemprichii S / m p h y l l i a radians Oxypora glabra Goniopora lobata A cropora n a s u t a A. ld . therefore a n n u a l g a m e t o g e n i c cycles are assumed. 2d . d..q ld. 27 Nov. l . l*~-I 1 " .. 1" I ld. 1983.I If ..R.ld. Harriott. d. le 2d ld t ld t lb.. grandis Faviafavus Favites h a # c o r a Leptoria phrygia O u l o p h / l l i a crispa Mycedium elephantotus A ustralogyra zelli A. 1' 1 lb lb lb lb lb lb lb.-[ 2b ~ . no. F o r the r e m a i n d e r o f the species involved. It is possible that the total n u m b e r o f species i n v o l v e d in these mass s p a w n i n g s is e v e n greater t h a n we h a v e so far r e c o r d e d . For further details see legend to Table 3 Species n No. florida F a v i a stelligera A cropora tenuis Montastrea magnistellata A cropora h y a c i n t h u s A . sarmentosa A . full moon was 20 November. p e c t i n a t a Platygyra lamellina P. ld. ld 2b. of species observed = 42. e i g h t e e n species r e c o r d e d in the mass s p a w n i n g h a v e b e e n r e p o r t e d to h a v e a n n u a l g a m e t o g e n i c cycles t e r m i n a t i n g in a b r i e f r e p r o d u c t i v e p e r i o d ( M a r s h a l l a n d S t e p h e n s o n . 2d. 1985). it is unlikely that they could h a v e spawned at o t h e r times o f the year. p e r s o n a l c o m m u n i c a t i o n ) . 2 $ - - . In a d d i t i o n to the fifteen species w h o s e g a m e t o g e n i c cycles w e r e m o n i t o r e d in the p r e s e n t study.l d . Favites a b d i t a Astreopora myriophthalma Goniastrea retiformis P l a t y g y r a daedalea A cropora cerealis A .lb. ld. 2d . ld 2b f t 4b - I - (partial) t ld. d. or o v e r 50% o f the species listed by D o n e (1982) as c o n s t a n t or f r e q u e n t in coral c o m m u n i ties across the central G B R . formosa A. P r o x i m a t e factors i m p l i c a t e d in s y n c h r o n o u s s p a w n i n g The synchrony. 6"-------t ld lb.. 1982 a. Veron. a n d p e r s o n a l c o m m u n i c a t i o n . l*--J 2b 2b lb species w h i c h o c c u r on the G r e a t Barrier R e e f (J.! lb. rigida A cropora m i l l e p o r a A. humilis Favia matthai F. 5d---I • --lb. 3d* .: Synchronous spawnings of scleractinian corals Table 7. of species for which direct spawning observations were made = 27. 1"-t -lb.

5F 4. Day column gives the day after the full moon. stelligera Favites abdita F. 6F 4. F: field observation.C.4F 3. 5F 5F 4. 6A 2:10 0:40 4. 6F 4F 1:50 1:55 3:20 3:10 0:25 2:20 2:55 3:40 3:10 2:20 1:10 3:10 5A 6A 4A 5A 4.formosa A. valida Astreopora myriophthalma Montipora digitata Agaricidae 4F 15A 3F 3F 3F 3F 2:30 2:45 1:55s 2:40 2:35 4F 3:40 2:45 5A 5A 7A 5A 6A 6. 2F 5. median values are given (negative sign indicates spawning before sunset). 7A 2:25s 0:10 2:55 1:50 0:10 1. 5. hour column shows hour:min after sunset at which spawning occurred. 7A 5A 6A 7A 5A 7A 5A 6A 4A 1:40 3:15 2:05 3:20 3:10 1:25 3:15 3:40 1:25 3:40 1:15 1:05 2:05 6F 6F 6F 6F 6F 6F 3:50s 3:40 3:50s 3:50s 3:10 6A 6F 6A 6A 6A 4A 4F 1:05 3:40 3F 3F 3F 1. The possible importance of temperature in determining timing of the spawning season is supported by the dif- . 6F 4. S: setting of egg-sperm bundles only. Species observed spawning at more than one reef in 1983. hyacinthus A. release not observed Species Magnetic day Acroporidae h Orpheus day h Lizard day h Bowden day h Big Broadhurst day h Acropora cerealis A. 5A 5A 5A 2:55 l:15 6F 2:25 5A 5. favulus G. nasuta A. monthly lunar or tidal cycles. A: aquarium observation. and diel light cycles. Mass spawning occurs in the period between midOctober and early December after a rapid spring rise in sea temperature between late August and September (Fig. sinensis Merulinidae 5A 4F 6A 6A 3. halicora Goniastrea aspera G.390 R. 5F 1:00 6F 6F 3:40 3:50s 6A 6A 6F 2:25 6A 5A 6A 2:10 2:40 Pachyseris speciosa Faviidae A ustralogyra zelli Cyphastrea chalcidicum Favia pallida F. 6F 3:25 3:25 Goniophora lobata cues which operate on increasingly fine time scales: a n n u a l sea temperature patterns. 5A 5A 5A 6A 5A 4. retiformis Leptoria phrygia Montastrea magnistellata Oulophyllia crispa Platygyra daedalea P. 5. nobilis A.: Synchronous spawnings of scleractinian corals Table 8. 6F 2:40 Galaxea fascicuIaris Pectinidae Echinophyllia orpheensis Mycedium elephantotus Pofitidae 5F 5F 4. 6.5F 4F 5. 5F 4. 7A Merulina arnpliata Oculinidae 5. secale A. 5A 5A 5A 4F 1:25 4. humilis A. 2F 5. 5). millepora A. 4F 1:55 4F 1:50 1:15 3:25 1:10 4. tenuis A. Where more than one colony was observed. Sea water temperatures along the G B R show a distinct seasonal variation which m a y influence gametogenic cycles.05 to 2:30 3:40 3:20 0:40 3:10s 5F 3:40 4:20s 4:10 4:30 2:20 2:55 3:40 5:00 3:10 l:35 1:20 3:25 1:30 1:45 5A 6A 5F 6. Babcock et al. 5F -0. 7A 5A 4:10 2:10 4:50 6A 4F 3F 4F 1:15 1:25 3:25 3:45 5F 3:40 6. elseyi A.

Most species spawned after the full moon and two species spawned after the new moon (Fig. A wide range of other marine organisms also display distinct lunar periodicity in their reproductive behaviour (Korringa. 1984) are similar to many corals in that they occur on only a few days once a year. Average annual temperature variation at Magnetic Island during 1979-1983 and Palm Islands (Orpheus Island and Eclipse Island) during 1979-1984. == =< 25 20 J F M A M J J A S O N D MONTH Fig.. the spawnings of Cornanthus japonica (Kubota. C. Acropora elseyi. Similar mech- .R. Temperatures were plotted and curves fitted by eye ferences in the month of spawning at the inshore and offshore reefs.. 1984. 1937. 1979. 1984) and in gamete-releasing species (Kojis and Quinn 1981.PALM ISLANDS . 1985). with Magnetic Island populations spawning one month earlier than those at offshore reefs (Big Broadhurst. 1981). 1982a. This does not prove the existence of an endogenous lunar rhythm.. Korringa.. 1981). 1981) and Eunice viridis (Caspers.. It has been shown that the timing of the release of planulae in races of PocilIopora damicornis at Hawaii can be manipulated by means of artificial moonlight (Jokiel et al. The same pattern was observed in 1983. ~ ... Kubota. In particular. Richmond and Jokiel. 6). Only one species. 1: Babcock (1984). Harrison etal. b. in early December (Harrison et al. 1984). Annual temperature variation at Magnetic Island 1980 1983. 6. 3: present study ~ 30' "*'. M1 the corals recorded in this paper exhibited lunar periodicity in reproductive behaviour. but indicates that moonlight is an entraining factor. recorded at 2 to 5 m depth.. Lizard and Orpheus). Babcock (1984) showed that for Goniastrea aspera in aquaria. Arrows indicate observed spawning periods. 1947.. It appears likely that the final sequence of events leading to the release of gametes is set in motion by the onset of darkness. Babcock et al. 1953. recorded at 3 to 5 m depth. 1920. it occurred one lunar month later. 2). Babcock. Other factors such as pressure (Naylor and Atkinson. 1984). Atoda. Sea temperatures began to rise earlier and more rapidly in the shallow (maximum depth 10m) inshore waters around Magnetic Island than they did further offshore at Orpheus Island in the Palm Island Group (Fig. 2: Harrison et al.. while at Big Broadhurst Reef and Orpheus Island.. Lunar periodicity has been well documented in brooding corms (Abe.. once the gametes are fully differentiated. Spawning tends to occur after a specific period of darkness. Changes in temperature have previously been associated with annual periodicity of spawning in other marine in- vertebrates (Orton. Our observations of mass spawning in corals suggest that the release of gametes occurs after nightfall following the first full moon subsequent to the maturation of gonads. ranging from a few minutes to over four hours after sunset (Tables 1 and 8).. 1974. Harriott. 1957. spawning at Magnetic Island took place early in November. (1984). Grigg. Rinkevich and Loya. Johannes. spawned at both the new and full moons. 1972) and water motion (Neumann. but they have not been demonstrated in coelenterates. 1978) may act as "Zeitgebers" for a tidal rhythm. 5. Campbell. Lewis.: Synchronous spawnings of scleractinian corals 391 30 o 25 20 J FM A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M AM 1980 1981 1982 J J A S ON 1983 D J F Fig. In 1982. the hour of spawning could be controlled by manipulating the light-dark cycle. Bowden. 1983. 1974. 1979) as well as scleractinians (Kojis and Quinn. MAGNETICISLAND ' SPAWNING PERIOD . This earlier warming may accelerate the maturation of gonads. 1978.

therefore. spawning occurs around low tide. The significance of the precise time during the lunar/tidal cycle at which coral spawning occurs may be related to the need to avoid predation while still producing a high concentration of gametes in order to maximize fertilization. The interspecific synchrony of coral spawning on the GBR is the most significant and unusual aspect of the mass spawning phenomenon.: Synchronous spawnings of scleractinian corals Evidence which supports the importance of low tides as a factor in determining the timing of spawning can be obtained from Goniastreafavulus. moonlight and sunlight may be used by corals as cues to synchronize spawning.(~. This activity was observed only during the crepuscular period. R. and larval development (Heyward and Babcock. 1968). which picked off egg-sperm bundles as they were released.C. and Pomacentrus. 1984) that the chances of survival of planktonic larvae would be increased during synchronous multispecific spawning by satiating predators and filter-feeders. they are not necessarily the forcing functions which ultimately determine the time when spawning will take place. moonlight can be used to predict the tidal regime.. however it is achieved. including species of Abudefduf Neopomacentrus. Spawning at night minimizes predation by visual feeders such as planktivorous fish species. during which the majority of coral species were observed to spawn. This species spawns during daylight (16. 7. A number of planktivorous fish species have been observed eating coral eggs. the physical constraints which determine optimal spawning times for each species are also likely to be similar. but after dark (18. Thus. While temperature. Yoshida et al. Tidal cycle. . 1980.5 m).. resulting in an extended period of slack water (Fig. October and November 1983 for Townsville area. This use of one signal to gain information on another. 7). The difference between this low tide and the following high tide is small (0..00-18. (Campbell. maximizes the probability of achieving fertilization. 1981). Hoffman. 1978).25-20. Triangles indicate days on which corals were known to have spawned. 1962. 1980). Tides in the Townsville region always occur one half hour or more later than those at Heron (Maxwell.iI. 1985) of most of the species involved in the mass spawning are very similar. . modes of reproduction.. it is possible that the timing of spawning is ultimately due to the advantage of increased fertilization during periods of low water motion and low water volume. when there was still sufficient fight for Possible causal factors of synchronous spawning Successful synchronization of gamete release within populations. Tide heights themselves appear not to trigger the final events leading to spawning since corals kept in aquaria still spawn at the appropriate time as determined by the light dark cycle.392 anisms using diel changes in light appear to be widespread in coelenterates. On the nights following the full moon. perhaps less easily detectable factor. It has also been suggested (Harrison et al. . 1984).U I- 3 2 a_ 1 0 • OCTOBER NOVEMBER Fig. 1974. has been shown to exist in a number of arthropod species (Tevis and Newell. This synchrony may be the result of species responding independently to physical factors in order to maximize reproductive success.Q. In both cases. For example. however. Honnegger et al. . The buoyant properties of egg-sperm bundles. MOON PHASE . • • "0 4 I ° "" "• "O" " I. Numbers beneath triangles indicate number of species from all five reefs which were known to have spawned on those days . 1969). ..50hrs) on reefs near Townsville.00hrs) at Heron Island (Kojis and Quinn. It is also possible that the timing of spawning is related to the advantage of reduced dispersal of gametes prior to fertilization.U -r" I. Babcock et al.. low tide falls between late afternoon and midnight on the GBR. . Spawning during low tide has been postulated as a mechanism for increasing the concentration of gametes in other animals (McDowall. even when that cycle is artificially manipulated (Babcock.

: Pink stripe on the ocean. 65-84 (1978) Jokiel. C. and the Great Barrier Reef Marine Park Authority. J. Quinn: Reproductive ecology of two faviid corals (Coelenterata. some corals appear to spawn together (Acropora spp.: Reproductive ecology of four scleractinian species at Lizard Island. Science. A. R. Ed. Freshwat. A. Part 1. E. J.R. 219-245 (1933) Maxwell.S. the Australian Coral Reef Society. This research was supported by Commonwealth Post-Graduate Awards and grants from M. Qninn: Aspects of sexual reproduction and larval development in the shallow water hermatypic coral Goniastrea australiensis (=favulus) (Edwards and Haime. 2. Coral Reefs 1. Am. Z. 8. Aust. B) 95. 36. Collins: Growth and sexual reproduction in the scleractinian coral Montipora digitata Dana. University of the Phillipines) Kojis. Veron for the identification of corals. Coral Reefs 2. Rep. Phillipines: Marine Sciences Center. Gomez et al. C. Mellors. J.: Lunar periodicity in reproduction.: Postlarval development of the coral Fungia actiniformis var. Gt Barrier Reef Exped. pp 69-74. Liu: Night irradiance and synchronization of lunar release of planula larvae in the reef coral Pocillopora damicornis. E. Littlefield. Pichon and J. R. Quezon City.: Spawning periodicity and habitat of the palolo worm Eunice viridis (Polychaeta: Eunicidae) in the Samoan Islands. The corals. mar. R. It may be significant that the majority o f corals spawn during the period of darkness between sunset and moonrise.G. 88. S. Mar. Bowden. Alderslade. J. R. Biol. 258 pp. J. G. but a greater number spawn or planulate at various times throughout the year (Szmant-Froelich etal. and A. D. coral Reefs 2. Elsevier 1968 McDowall. Tardent: Light controlled spawning in Phialidium hemisphaericum (Leptomedusae). L. Trench and J. J. and T. by A. 89. 105-121 (1953) Babcock. C. Scleractinia). 5th int.. and T. 1985). pp 83-88. M. R. and N. A.. Harrison. Baenninger and P. Ed. French Polynesia: Antenne Museum-EPHE) Heyward. (Ser. Gabrie et al. D. Lovell.: Reproduction and distribution of two species of Goniastrea (Scleractinia) from the Great Barrier Reef Province. H. P. P. Bull. Tuatara 17. (Ed. D. by E. New York: Elsevier/North Holland 1981 Lewis.. 1186-1189 (1984) Heyward. Symp. A. Florian for assistance in photography. Slicks of coelenterate larvae have been reported from Bermuda (Butler. In: Reproductive ecology of marine invertebrates. 441-446 (1985) Hoffman. on the third to sixth nights after full moon. M. and N. Stephenson: The breeding of reef animals. Jackson. p. by W. mar. Sorokin. Adams. J.. Mar.: Reproductive ecology of two species of gorgonian corals: relations to vertical and geographical distribution. 133-144 (1969) Literature cited Abe. B. Great Barrier Reef. Willis for supplying us with data on the spawning of soft corals. M. R.: Reproductive strategies of coastal marine fishes in the tropics. Palao trop. Y. Biol. 187-195 (1984) Bothwell. New York: Academic Press 1974 . R. Scient. Nat. 1984). V. B. coral Reefs 2. Veron. 62-63 (1980) Campbell. L. E.: Synchronization of spawning in the crinoid. Monogr. It is not yet known whether the mass spawning of corals observed on the G B R occurs in other parts of the world. In the Caribbean Sea. 15. Tardent. Stoddart. 1980).. 167-174 (1985) Kojis.: Cnidaria. Ecol. 167-173 (1974) Marshall. 917-933 (1957) Kubota. We would like to thank the following: P. R. G. mar. where coral species spawn over a period of several months. J. N. 523-550 (1924) Grigg. J. 347-381 (1947) Korringa. geol. H. A general review. appears to be more pronounced on the G B R (Maxwell. and R. 1968) than in the Red Sea (Shlesinger and Loya. 3. pp 133-139. L. Pearse. The phenomenon does not appear to occur in the Red Sea. Soc. R. W. P. 137-144 (1982). and N. D. S. Proc. Sci.: The larvae and post larval development of the reef building coral. 67. N. N. Babcock. Coral Reefs 2. In: Reproduction in marine invertebrates. Clark Jr. 145-152 (1982a). Veron for providing comments on the manuscript. P. G. Agostino. Proc. R. 3I. 1. Ito and P. and video camera work. Tapiolas. J. Nat. Wallace and B.: Environmental uncertainty and evolution of physiological adaptation in Colias butterflies. J.: Lunar rhythms in aquatic animals. N. B. Ecol. D. Ed. 4th int. Am. pp 41-59. 393 Caspers. Watt for field assistance. K. Quezon City.S. H. L.Y. J. 1980) which appear to be similar to slicks observed as a result o f coral spawning on the GBR. D.Y. (Ser. T. Symp. by P. K. D. Prog. a means of asexual reproduction and dispersal in the coral genus Acropora (Scleractinia: Astrocoeniida: Acroporidae) . In: Developmental and cellular biology of coelenterates. Ed. R.: Synchronous spawnings of scleractinian corals visual feeding. Ecol. J. 79. 9-18 (1983) Harrison. P.: Relationship between the moon and periodicity in the breeding of marine animals. New York. Ayre. 1857). J. thereby concentrating the reproductive season o f m a n y corals into a relatively brief period o f time. 17. 112. W. 4) 20. Babcock. Mere. Collins. 223. Giese and J. Mar. C. J. L. by C. Gomez et al. C. and during a variety of different lunar phases (Shlesinger and Loya. which may determine the most favourable spawning period. Stn Stud. Sci. Harriott. B. H.. 1L C. 251-255 (1982b) Korringa. Stidwill. Pini. J. Babcock: Embryonic and post embryonic development of some hermatypic scleractinians. Phillipines: Marine Sciences Center. 4th int. Moore. 1. 558-573 (1981) Kojis. Res. such as tidal amplitude and annual sea temperature range.: Patterns in the distribution of coral communities across the central Great Barrier Reef. Proc. T. Hist. Biol. Achermann. Biol. S. Variation in physical factors. L. and J.a preliminary report.: Fragmentation. Stanyck. and R. Acknowledgements. No.: Settlement behaviour of the planulae larvae of the hermatypic coral Faviafragum (Esper). Vol. Ser. L. New York: Elsevier/North Holland 1980 Johannes. Bull. H. 176 (1985). Babcock et al. S. University of the Phillipines) Butler. palawensis Doderlein. Proc. J. Soc. Envir. V. This m a y more sharply define the period of time during which conditions are optimal for reproductive success on the GBR. 999-1015 (1978) Honnegger. Trhuku Univ. or Caribbean Sea (Olhorst. L. 229-236 (1984) Done. 3. 1985). In: Advances in invertebrate reproduction. Tardent and R. B. C. by E. B. Columbia: University of South Carolina Press 1979 Harriott. 9. by S.). M. Comanthus japonica.: Lunar periodicity. M. Fish. Biol.: Atlas of the Great Barrier Reef.T. microscopy. N. 95-107 (1982) Fox. Oliver. M. D. Willis: Mass spawning in tropical reef corals. (Ed. (Ed. A. coral Reef Congr. Moorea. 73-79 (1937) Atoda. exp. Rep. Quinn: Reproductive strategies in four species of Porites (Scleractinia).

: Porifera: sudden sperm release by tropical Demospongiae. Ecol. Synchronization in breeding and seasonality of planulae shedding. thesis. Ikegami: Darkness induced maturation and spawning in Spirocodon saltatrix. C.Y. Inst. by P. J. Ecol. A. Science. H. breeding and distribution in marine animals. and Y. W.C. Speciesspecificity and sperm behavior. Science. S. Jr. N.Y. E. recruitment and fragmentation in nine sympatric species of the coral genus A cropora. R. L. H. Y. S. Mar. mar. mar. J. pp 75-82. II. 80 pp. exp. J.: Sea temperature. J. 538-539 (1970) Richmond. Ed.D. 88. 1985. mar. M. Tardent and R. Biol. M. Pichon: Scleractinia of eastern Australia. biol. H. Loya: Coral community reproductive patterns: Red Sea versus the Great Barrier Reef. Ser. 395-415 (1972) Neumann. 12. 1-159 (1982) Wallace.: Jamaican coral reefs: important biological and physical parameters. 280-287 (1984) Rinkevich. Atkinson: Pressure and the rhythmic behaviour of inshore marine animals.: Synchronization of reproduction in marine insects by tides. Hon.: Intra and interspecific interactions of two species of Montipora. Mar. Bull. Ecology 43. and R. thesis. J. Biol. Jr. 23. Adams. 1. Soc. A. L. Humphrey. 145-152 (1979) R.394 Miller. Aust. P. 1. 34. N.. F. 339-366 (1920) Reiswig. and M. by W. ln: Advances in invertebrate reproduction. pp 21-35. 53.. Prog. 163 pp. 497-505 (1962) Veron.: Synchronous spawnings of scleractinian corals Robertson. 217-233 (1985) Yoshida. Part IV: family Acroporidae. Newell: Studies on the biology and seasonal cycle of the giant red velvet mite. New York: Elsevier/North Holland 1980 Date of final manuscript acceptance: September 6. Sci. Monogr. J. 99-114 (1979) Naylor.K. Ph. U. Ed. A. Ser. New York: Elsevier/ North Holland 1981 Olhorst. N. R. C. Tardent. Communicated by G. and Y. N. 35. exp. mar. Sydney . Honji and S. Sci. and I. Riggs and M. Jokiel: Lunar periodicity in larva release in the reef coral Pocillopora damicornis at Enewetak and Hawaii. Am. L. Loya: The reproduction of the Red Sea coral Stylophora pistillata. 961 (1984) Tevis. Zool. B. M. Biol. p. 1333-1335 (1985) Szmant-Froelich. J.: Entrainment of a semilunar rhythm by simulated tidal cycles of mechanical disturbance. Clark. Dinothrombium pandorae (Acari: Thrombiidae). 73-85 (1978) Neumann. James Cook University of North Queensland 1981 Shlesinger. Babcock et al. L. 170. 26.: Sperm chemotaxis in the hydromedusae. Yale University 1980 Orton. 5. Biol.: Reproduction. and T. E. Reutter: Sexual reproduction in Caribbean reef corals. D. In: Developmental and cellular biology of coelenterates. and P. Mar. Symp. Ass. 228. H.